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      A Potential Role for Bat Tail Membranes in Flight Control


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          Wind tunnel tests conducted on a model based on the long-eared bat Plecotus auritus indicated that the positioning of the tail membrane (uropatagium) can significantly influence flight control. Adjusting tail position by increasing the angle of the legs ventrally relative to the body has a two-fold effect; increasing leg-induced wing camber (i.e., locally increased camber of the inner wing surface) and increasing the angle of attack of the tail membrane. We also used our model to examine the effects of flying with and without a tail membrane. For the bat model with a tail membrane increasing leg angle increased the lift, drag and pitching moment (nose-down) produced. However, removing the tail membrane significantly reduced the change in pitching moment with increasing leg angle, but it had no significant effect on the level of lift produced. The drag on the model also significantly increased with the removal of the tail membrane. The tail membrane, therefore, is potentially important for controlling the level of pitching moment produced by bats and an aid to flight control, specifically improving agility and manoeuvrability. Although the tail of bats is different from that of birds, in that it is only divided from the wings by the legs, it nonetheless, may, in addition to its prey capturing function, fulfil a similar role in aiding flight control.

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          Most cited references80

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          Flying and swimming animals cruise at a Strouhal number tuned for high power efficiency.

          Dimensionless numbers are important in biomechanics because their constancy can imply dynamic similarity between systems, despite possible differences in medium or scale. A dimensionless parameter that describes the tail or wing kinematics of swimming and flying animals is the Strouhal number, St = fA/U, which divides stroke frequency (f) and amplitude (A) by forward speed (U). St is known to govern a well-defined series of vortex growth and shedding regimes for airfoils undergoing pitching and heaving motions. Propulsive efficiency is high over a narrow range of St and usually peaks within the interval 0.2 < St < 0.4 (refs 3-8). Because natural selection is likely to tune animals for high propulsive efficiency, we expect it to constrain the range of St that animals use. This seems to be true for dolphins, sharks and bony fish, which swim at 0.2 < St < 0.4. Here we show that birds, bats and insects also converge on the same narrow range of St, but only when cruising. Tuning cruise kinematics to optimize St therefore seems to be a general principle of oscillatory lift-based propulsion.
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            Bat flight generates complex aerodynamic tracks.

            The flapping flight of animals generates an aerodynamic footprint as a time-varying vortex wake in which the rate of momentum change represents the aerodynamic force. We showed that the wakes of a small bat species differ from those of birds in some important respects. In our bats, each wing generated its own vortex loop. Also, at moderate and high flight speeds, the circulation on the outer (hand) wing and the arm wing differed in sign during the upstroke, resulting in negative lift on the hand wing and positive lift on the arm wing. Our interpretations of the unsteady aerodynamic performance and function of membranous-winged, flapping flight should change modeling strategies for the study of equivalent natural and engineered flying devices.
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              Animal flight dynamics I. Stability in gliding flight.

              Stability is as essential to flying as lift itself, but previous discussions of how flying animals maintain stability have been limited in both number and scope. By developing the pitching moment equations for gliding animals and by discussing potential sources of roll and yaw stability, we consider the various sources of static stability used by gliding animals. We find that gliding animals differ markedly from aircraft in how they maintain stability. In particular, the pendulum stability provided when the centre of gravity lies below the wings is a much more important source of stability in flying animals than in most conventional aircraft. Drag-based stability also appears to be important for many gliding animals, whereas in aircraft, drag is usually kept to a minimum. One unexpected consequence of these differences is that the golden measure of static pitching stability in aircraft--the static margin--can only strictly be applied to flying animals if the equilibrium angle of attack is specified. We also derive several rules of thumb by which stable fliers can be identified. Stable fliers are expected to exhibit one or more of the following features: (1) Wings that are swept forward in slow flight. (2) Wings that are twisted down at the tips when swept back (wash-out) and twisted up at the tips when swept forwards (wash-in). (3) Additional lifting surfaces (canard, hindwings or a tail) inclined nose-up to the main wing if they lie forward of it, and nose-down if they lie behind it (longitudinal dihedral). Each of these predictions is directional--the opposite is expected to apply in unstable animals. In addition, animals with reduced stability are expected to display direct flight patterns in turbulent conditions, in contrast to the erratic flight patterns predicted for stable animals, in which large restoring forces are generated. Using these predictions, we find that flying animals possess a far higher degree of inherent stability than has generally been recognized. This conclusion is reinforced by measurements of the relative positions of the centres of gravity and lift in birds, which suggest that the wings alone may be sufficient to provide longitudinal static stability. Birds may therefore resemble tailless aircraft more closely than conventional aircraft with a tailplane.

                Author and article information

                Role: Editor
                PLoS One
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                30 March 2011
                : 6
                : 3
                : e18214
                [1 ]Faculty of Life Sciences, University of Manchester, Manchester, United Kingom
                [2 ]Département d'aérospatiale et mécanique, Université de Liège, Liège, Belgium
                University of Western Ontario, Canada
                Author notes

                Conceived and designed the experiments: JDG GD. Performed the experiments: JDG GD. Analyzed the data: JDG RLN. Contributed reagents/materials/analysis tools: GD JRC. Wrote the paper: JDG RLN GD JRC.

                Gardiner et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
                : 22 September 2010
                : 28 February 2011
                Page count
                Pages: 8
                Research Article
                Anatomy and Physiology
                Comparative Anatomy
                Evolutionary Biology
                Organismal Evolution
                Animal Evolution
                Comparative Anatomy
                Aerospace Engineering
                Fluid Mechanics



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